Review and identification of the cuckoo bees of central Europe (Hymenoptera: Halictidae: Sphecodes)

Accepted by C. Rasmussen: 20 Jan. 2012; published: 14 May 2012

ZOOTAXAISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)Copyright © 2012 · Magnolia Press

Zootaxa 3311: 1–41 (2012) www.mapress.com/zootaxa/ Article

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Review and identification of the cuckoo bees of central Europe (Hymenoptera: Halictidae: Sphecodes)

PETR BOGUSCH1 & JAKUB STRAKA2

1University of Hradec Králové, Department of Biology, Rokitanského 62, CZ-500 03 Hradec Králové, Czech Republic. E-mail: [email protected] University in Prague, Faculty of Science, Department of Zoology, Viničná 7, CZ-128 44 Praha 2, Czech Republic. E-mail: [email protected]

Table of contents

Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3Identification key of Sphecodes of central Europe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3List of species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

Sphecodes albilabris (Fabricius, 1793) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7Sphecodes alternatus Smith, 1853 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8Sphecodes crassanus Warncke, 1992 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8Sphecodes crassus Thomson, 1870. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9Sphecodes cristatus Hagens, 1882 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9Sphecodes croaticus Meyer, 1922 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9Sphecodes dusmeti Blüthgen, 1924 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10Sphecodes ephippius (Linné, 1767) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10Sphecodes ferruginatus Hagens, 1882 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10Sphecodes geoffrellus (Kirby, 1802) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10Sphecodes gibbus (Linnaeus, 1758) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11Sphecodes hyalinatus Hagens, 1882 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11Sphecodes intermedius Blüthgen, 1923 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12Sphecodes longulus Hagens, 1882 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12Sphecodes majalis Pérez, 1903 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12Sphecodes marginatus Hagens, 1882 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13Sphecodes miniatus Hagens, 1882 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13Sphecodes monilicornis (Kirby, 1802) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13Sphecodes niger Hagens, 1874 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14Sphecodes nomioidis Pesenko, 1979. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14Sphecodes olivieri Lepeletier, 1825 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14Sphecodes pellucidus Smith, 1845 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15Sphecodes pinguiculus Pérez, 1903 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15Sphecodes pseudofasciatus Blüthgen, 1925 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16Sphecodes puncticeps Thomson, 1870 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16Sphecodes reticulatus Thomson, 1870 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16Sphecodes rubicundus Hagens, 1875 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17Sphecodes ruficrus (Erichson, 1835) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17Sphecodes rufiventris (Panzer, 1798) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17Sphecodes scabricollis Wesmael, 1835. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17Sphecodes schenckii Hagens, 1882 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18Sphecodes spinulosus Hagens, 1875 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18Sphecodes zangherii Noskiewicz, 1931 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

Acknowledgement . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39

TERMS OF USEThis pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.

BOGUSCH & STRAKA2 · Zootaxa 3311 © 2012 Magnolia Press

Abstract

We reviewed nomenclature, biology, hosts, geographical distribution and compiled an identification key for all 33 Sphe-codes Latreille, 1804 species known from central Europe. The identification key is separated for females and males andinclude 204 figures (photographs) of identification characters as well as male genitalia of all species. Taxonomically dif-ficult groups within the genus were critically studied and new characters, as well as corrected geographical distribution,are presented, i.e., the S. reticulatus group (S. alternatus Smith, 1853, S. crassanus Warncke, 1992 and S. reticulatusThomson, 1870), S. croaticus group (S. croaticus Meyer, 1922, S. pseudofasciatus Blüthgen, 1925 and S. zangherii Nosk-iewicz, 1931) and S. miniatus group (S. marginatus Hagens, 1882, S. miniatus Hagens, 1882 and S. nomioidis Pesenko,1979). The name S. nomioidis is used because it is the only available name for the taxon formerly identified as S. margi-natus in Eastern Europe. Sphecodes capverdensis Pesenko & La Roche, 2002 is considered to be a junior synonym of S.pinguiculus Pérez, 1903 (syn. nov.). In addition we summarized all known host records of Sphecodes, including a discus-sion of the likelihood of published data and presentation of new host data.

Key words: identification key, taxonomy, ecology, hosts

Introduction

The bees (Hymenoptera: Apoidea: Apiformes) represent one of the richest groups of aculeate Hymenoptera interms of species. Yet, even as of recently several taxonomicaly complicated groups remain poorly studied andrarely reviewed. Such typical groups are within the family Halictidae where the only European cuckoo bee genus,Sphecodes Latreille, 1804, has been studied several times, but remains one of the most difficult European bee gen-era to identify at the species level. Knowledge on the taxonomy of the genus was summarized by Hagens (1874,1875, 1882), who also described the majority of all currently valid species. His descriptions were for the most partthe source of information for compilation of later identification keys. Additional species were described by Meyer(1919, 1922) and Noskiewicz (1931). Recently, eight new species from Italy were described by Campadelli &Nobile (2000) and Nobile & Turrisi (2004). However, the validity of these species is doubtful (Schwarz & Gusen-leitner 2012) and thus, we do not include them in this article. Burger & Reum (2004) and Burger et al. (2006)examined difficult groups of this genus, in particular S. croaticus Meyer, 1922 and S. zangherii Noskiewicz, 1931.European Sphecodes have been keyed four times. Blüthgen (1923a) compiled the first key, followed by Šustera(1959). Although the nomenclature used in those papers is old, the authors found many previously unknown char-acters. A useful key was published by Warncke (1992). This publication contains good drawings of male genitaliaand distribution maps of all European species. However, most of the key couplets are based on a single and some-times a very variable character. Thus the key is good but some species are hardly identifiable. The latest key, pub-lished by Amiet et al. (1999), is in comparison to Warncke’s key with fewer species but rely on better identificationcharacters.

The biology of Sphecodes was studied by few authors. Brief references are found in monographs on the bees ofEurope (Friese 1898, Stoeckhert 1930, Westrich 1989, Macek et al. 2010), where host species of Sphecodes arementioned occasionally. Blüthgen (1923b, 1934) provided the first review on the biology of Sphecodes, includingthe hosts. These data were cited by Westrich (1989), Celary (1991) and Amiet et al. (1999). Hosts of Sphecodeswere also studied by Vegter (1985, 1993). Sick et al. (1994) studied hosts of selected Sphecodes species experimen-tally and reported new information. Except some ecological studies reporting Sphecodes hosts and mentioned inthe text under the appropriate species, the following species have been studied in detail: S. cristatus Hagens, 1882in Sweden by Svensson (1982), S. majalis Pérez, 1903 by Herrmann et al. (2003), and S. ruficrus Erichson, 1852by Herrmann (2006). Host specificity of two common species, S. ephippius (Linnaeus, 1767) and S. monilicornis(Kirby, 1802) was studied by Bogusch et al. (2006). Data on the visited flowers were published by Westrich (1989)and Celary (1991), demonstrating polylectic behavior of all species. Sphecodes bees are nest cleptoparasites, theyonly forage on nectar on flowers and do not collect pollen. Thus the visited flowers are not discussed in this publi-cation.

Here, we would like to present a newly compiled identification key based on the study of the material of allincluded species, and emphasizing the advantages and eliminating the disadvantages of previously published keys.The publication also reviews the synonymy, taxonomy, distribution, and biology of all species included. Host spec-trum of all species is critically analyzed. The key includes all species known from central Europe and closely alliedareas.


Zootaxa 3311 © 2012 Magnolia Press · 3REVIEW OF THE CUCKOO BEES OF CENTRAL EUROPE

Methods

The nomenclature was adopted from the following sources: Warncke (1992), Schwarz et al. (1996) and Bogusch etal. (2007). The material studied is from the following collections: BMNH—The Natural History Museum, London,UK, HNHM—Magyar Természettudományi Múzeum Allattara, Budapest, Hungary, JSPC—Jakub Straka, privatecollection, Prague, Czech Republic, MSAC—Maximilian Schwarz, private collection, Ansfelden, Austria,NMPC—National Museum, Prague, Czech Republic, PBHK—Petr Bogusch, private collection, Hradec Králové,Czech Republic. Frank Burger (Weimar, Germany), Mike Herrmann (Konstanz, Germany), Toshko Ljubomirov(Sofia, Bulgaria), Denis Michez (Mons, Belgium) and Guido Pagliano (Torino, Italy) sent us gift material of Sphe-codes. In the key, the following abbreviations are used: T—tergite, S—sternite (T4 means the fourth tergite). Othermorphological terms are used in English and Latin languages. The characters in all parts are sorted from the mostimportant (best seen and least variable). Most of the characters were photographed and are shown under appropri-ate number within the captions. Numbers of figures are put in square brackets to the text of the key. The key alsocontains photos of male genitalia of all species from dorsal and lateral view. The list of species includes all speciesmentioned in the key, with the following information: name, synonymy, distribution and ecology (with a focus tohost species and biotope preferences). Only names and pages within the publication with description of all taxa arementioned, whole citations of the descriptions are in the reference list.

Identification key of Sphecodes of central Europe

Females (♀♀)

1 a. Clypeus about three times wider than long, with furrow in the middle and teeth on sides, shorter than supraclypeus [fig. 1];scutum finely and densely punctate, 6–8 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. majalis Pérez

1 b. Clypeus less than three times (usually 2x) wider than long, without teeth on sides [fig. 32]; scutum coarsely punctate . . . . . . 22 a. Vertex behind the ocelli with sharp transverse ridge [fig. 2]; flagellomeres one and half longer than wide, so the antenna looks

longer than other species; 1st flagellomere distinctly wider than 2nd [fig. 3], 8–11 mm . . . . . . . . . . . . . . . . . spinulosus Hagens2 b. Vertex without ridge; flagellomeres maximally as long as wide, equal in width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 a. Hind wing with 7–14 hamuli [fig. 4]; angle between cubital and anal veins on hind wing usually acute [fig. 6], if the angle

looks right, then head square and large; vertex behind the ocelli usually gibbous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43 b. Hind wing with 5–6 hamuli [fig. 5] (only in some specimens of S. pellucidus, S. rubicundus and S. ruficrus up to 8 hamuli but

they all have right angle between veins on hindwing); usually right angle between cubital and anal veins on hind wing [fig. 7];vertex behind the ocelli not gibbous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

4 a. Gena on backside with sharp ridge [fig. 8], sharp or weak (S. scabricollis) elongate ridge behind ocelli . . . . . . . . . . . . . . . . . 54 b. Gena without ridge, ridge behind ocelli absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75 a. Face (clypeus and paraocular area), pronotum and scutellum with dense felt-like white pubescence; base of T1 very coarsely

punctate; thorax often reddish [fig. 10], 8–11 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . olivieri Lepeletier5 b. Face, pronotum and scutellum without felt-like pubescence; base of T1 unpunctate; thorax black. . . . . . . . . . . . . . . . . . . . . . 66 a. Vertex behind the ocelli with elongate ridge [fig. 11]; scutum coarsely but sparsely punctate; tibiae and tarsi reddish; smaller

and slenderer species, 6–8 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . cristatus Hagens6 b. Vertex behind the ocelli only with weak elongate ridge, which is in some cases very poor; scutum very densely and coarsely

punctate [fig. 9]; legs dark; larger, robust species, 9–12 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . scabricollis Wesmael7 a. Genae behind the eyes wide, vertex gibbous; genal width similar as the width of compound eye; head square-shaped [fig. 12];

angle between cubital and anal veins on hind wing acute but nearly right (about 80°), 7–10 mm . . . . . . monilicornis (Kirby)7 b. Genae narrow, narrower than the compound eye; head looks flat and smaller; angle between cubital and anal veins on hind

wing always acute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88 a. Mesopleura and propodeal sides regularly and finely ridged, ridges reach to hypoepimeral area and scutellum [fig. 15]; legs . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . rufiventris (Panzer)8 b. Mesopleurae grained without elongate ridges [fig. 14] (except ventral part of mesopleurae and propodeal sides with coarser

ridges in S. schenckii); legs with pale or dark hair . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99 a. Scutum coarsely and densely punctate, interspaces smaller than punctures [fig. 13]; tergites also coarsely and densely punctate;

wings dark; large and conspicuous species with red abdomen with dark apex (only last tergite completely black), 11–15 mm .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . albilabris (Fabricius)

9 b. Scutum coarsely but sparsely punctate with shiny and large interspaces among punctures; tergites usually conspicuouslypunctate, but not as in S. albilabris . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

10 a. Basis of T4 densely punctate [fig. 18]; vertex behind ocelli not elevated [fig. 16]; genae very narrow, getting narrower justbehind the compound eyes; pygidium shiny and as wide as first flagellomere . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

10 b. Basis of T4 sparsely punctate [fig. 19]; vertex behind ocelli strongly elevated [fig. 17]; genae wider; pygidium dull, distinctlynarrower than first flagellomere . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13



11 a. Apical depression of T4 without punctures, but finely wrinkled [fig. 18]; T1 finely and sparsely punctate; face coarsely anddensely punctate, dull, 7–10 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. reticulatus Thomson

11 b. Apical depression of T4 punctate without wrinkles; T1 coarsely punctate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1212 a. Tergites densely punctate but with distinct interspaces (well recognized on T4); end of T4 only with few fine punctures [fig.

20]; frons with distinct shiny interspaces among punctures [fig. 22], 8–11 mm . . . . . . . . . . . . . . . . . . . . . . . alternatus Smith12 b. Tergites coarsely and densely punctate; T4 with riddle-like punctures without distinct interspaces [fig. 21]; also the end of T4

finely punctate; frons dull, very densely punctate [fig. 23], 8–11 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . crassanus Warncke13 a. Scutum only with few coarse and deep punctures [fig. 24]; abdomen nearly unpunctate (but very variable punctures); hair on

scapus as long as width of scapus [fig. 26]; face without dense white pubescence, 7–13 mm . . . . . . . . . . . gibbus (Linnaeus)13 b. Scutum more densely punctate in the middle than on sides [fig. 25]; hair on scapus twice longer than width of scapus [fig. 27];

face usually densely whitish haired, 8–11 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . schenckii Hagens14 a. Tergites (also T1) densely and finely punctate [fig. 28]; wings hyaline, pterostigma yellow; face above antennal socket with

dense whitish pubescence [fig. 31] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1514 b. Tergites only sparsely punctate, T1 in many cases only with few punctures; wings not hyaline, usually slightly infumate; face

above antennal socket with longer sparse hair . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1615 a. Scutum finely and sparsely punctate [fig. 29]; propodeum with fine sculpture; completely red to dark with only few red marks,

5–7 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pinguiculus Pérez15 b. Scutum in the middle coarsely but sparsely punctate [fig. 30]; propodeum with grainy sculpture; dark with red abdomen, 5–8

mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . intermedius Blüthgen16 a. Mandibles without lateral tooth [fig. 32]; small species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1716 b. Mandibles with lateral tooth [fig. 33]; small to big species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1817 a. Head distinctly wider than long [fig. 32]; genae narrowing just behind the compound eyes; T3 densely and coarsely punctate

[fig. 37], 5–7 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . puncticeps Thomson17 b. Head only a little wider than long [fig. 34]; genae wider; T3 only sparsely punctate [fig. 38], 4–5 mm. . . . . longulus Hagens18 a. Hypoepimeral area smooth and shiny [fig. 35]; head nearly square-shaped, genae nearly as wide as compound eyes;

propodeum rugous only anteriorly [fig. 36]; all sterna reddish to brown, 4.5–5.5 mm . . . . . . . . . . . . . . . . . . . . . niger Hagens 18 b. Hypoepimeral area and the whole propodeum rugous; at least posterior sterna black; small to large species. . . . . . . . . . . . . . 1919 a. Pygidium wide and dull [fig. 39]; clypeus gibbous with medial furrow, all coarsely punctate [fig. 42]; scutum coarsely

punctate; medium size to big species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2019 b. Pygidium narrow and shiny, without sculpture [fig. 40]; clypeus only sparsely punctate; scutum finely punctate or with only

few coarse punctures [fig. 41]; small species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2320 a. Face and thorax with long black hair; tibiae and tarsi reddish; abdomen red only with black end (last tergite) [fig. 43], 6–9 mm

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ruficrus Erichson20 b. Face and thorax with short and pale hair, if with longer dark hair, then legs dark and/or abdomen with at least last 2 tergites

black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2121 a. T4 red on sides and dark in the middle; apical part of T1 delimited by two rows of punctures [fig. 45]; angle between the

cubital and anal veins on hind wing 90° or slightly more [fig. 44], 8–12 mm . . . . . . . . . . . . . . . . . . . . . . . rubicundus Hagens21 b. T4 completely dark; apical part of T1 delimited by a row of punctures [fig. 46]; angle between cubital and anal veins on hind

wing less than 90° . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2222 a. Head wider than long; clypeus with very long, pale hair; hair on scapus twice longer than width of scapu [fig. 47]s; inner side

of front tibiae always dark, 7–11 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pellucidus Thomson22 b. Head only a little wider than long; clypeus with shorter and sparse pubescence; hair on scapus as long as the width of scapus;

inner side of front tibiae usually (but not always!!) with reddish or yellowish spot, 6–9 mm . . . . . . . . . ephippius (Linnaeus)23 a. Pronotal projections round, not angulated [fig. 48]; dorsal part of propodeum round on sides, without sharp ledge [fig. 50]; face

very finely and densely punctate; clypeus flat without central furrow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2423 b. Pronotal projections with sharp angle [fig. 49]; dorsal part of propodeum on sides with sharp ledge; face more coarsely but

sparsely punctate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2524 a. Face around antennal sulcus dull, frons densely punctate [fig. 53]; ventral part of thorax ridged [fig. 51], 6–9 mm . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ferruginatus Hagens24 b. Face around antennal sulcus with shiny interspaces among punctures [fig. 54]; ventral part of thorax only finely sculptured

[fig. 52], 5–7 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . hyalinatus Hagens25 a. Face square-shaped (like in S. monilicornis), outer margin of eye straight; inner margins of compound eyes parallel (see from

above) [fig. 55]; genae strongly developed behind eyes [fig. 56] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2625 b. Face oval, outer margin of eyes largely rounded; inner margin of compound eye (in S. croaticus only a little) convergent [fig.

62] (see from above); genae narrower (except some S. croaticus). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2726 a. Eyes distinctly hairy (in old individuals can be worn); lower gena densely and uniformly hairy; interocellar distance smaller

than ocelloocular distance; occiput elevated above ocelli (distinct in frontal view); interocellar area with large interspacesamong punctures [fig. 57, 58]; posterolateral part of mesopleurae dull, at most with fine punctures; T1 finely and in second halfalso densely punctate, 5–8 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . zangherii Noskiewicz

26 b. Eyes at most with few short hair; lower gena sparsely hairy; ocelli on the top of occiput (distinct in frontal view); interocellardistance as long as ocelloocular distance; interocellar area densely punctate [fig. 55, 56]; posterolateral part of mesopleuraewith distinct bright shiny area with well-developed punctures; T1 finely and sparsely punctate, 5–7 mm . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pseudofasciatus Blüthgen

27 a. Furrow in the middle of frons extending at most to the middle of frons [fig. 59, 60]; T1 with diffuse punctures; T2 and T3



densely punctate in front half [fig. 61], 5–7 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . croaticus Meyer27 b. Furrow in the middle of frons almost touching front ocellus [fig. 74, 75]; tergites usually finely and sparsely punctate or with

diffuse punctures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2828 a. Scutum densely punctate; legs dark; supraclypeus very densely punctate [fig. 62], interspaces among punctures

microsculptured, dull, 4–6 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . dusmeti Blüthgen28 b. Scutum sparsely punctate; legs dark or pale; supraclypeus sparsely punctate, shiny . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2929 a. Flagellomere 3 longer than 1 and about as long as 4 [fig. 69]; T2 finely punctate or smooth; pygidium narrower [fig. 40], 4.5–

7 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3029 b. Flagellomeres 1, 2 and 3 similarly long, all distinctly shorter than long, no of them as long as flagellomere 4 [fig. 71]; at least

basal third of T2 and basal half of T3 with distinct punctures; pygidium wider [fig. 84], 4–6 mm . . . . . . (miniatus group) ...3130 a. Head distinctly wider than long [fig. 63]; flagellomere 3 longer than 2 [fig. 69]; scapus sparsely punctate ventromedially and

shiny; hypoepimeral area rugous [fig. 65]; T2 finely punctate [fig. 67], 5–7 mm . . . . . . . . . . . . . . . . . . . . . . crassus Thomson30 b. Head only slightly wider than long [fig. 64]; flagellomere 2 quadrate, wider and longer than flagellomere 3 [fig. 70]

(sometimes looks as subequal; measuring recommended); scapus densely punctate ventromedially; hypoepimeral area finelystriated [fig. 66]; T2 with a few punctures [fig. 68], 4.5–6 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . geoffrellus (Kirby)

31. Three species very hard to distinguish in females. No character fits in 100% of individuals: 31 a. Pygidial plate very wide, surface with distinct elevated longitudinal furrow in the middle [fig. 83, 84]; frons in front of the

oceli only slightly convex, densely punctate, distances among punctures variable and often form chains and elevated linesalong, punctures variable in size, some well-defined and round, some with irregular margins [fig. 74]; vertex distinct, fromlateral view slightly decreasing behind hind oceli [fig. 72]; T2 very finely punctate, punctures ill-defined; in the middle of T2,punctated area does not reach apical depression, puncures on the basal margin of apical depression distinctly isolated frompunctures of T2 base [fig. 80] (sometimes with more isolated punctures in this area, but punctures becoming posteriorly moreshallow and finer) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . miniatus Hagens

31 b. Pygidial plate usually narrowing posteriorly, or all narrower, surface flat, or indistinctly elevated in the middle [fig. 85, 86](when pygidium wider, than still indistinctly elevated in the middle); frons in front of the oceli strongly convex; frons sparsely,or densely, but very uniformly punctate, interspaces among punctures flat, most punctures uniform in size, well defined–round[fig. 76]; vertex short, from lateral view strongly decreasing immediately behind hind oceli [fig. 73]; T2 distinctly punctate,punctures small, but well-defined; in some specimens in the middle of T2 [fig. 81], punctation almost reach apical depressionor fully joining punctures on base of depression .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32

32 a. All frons uniformly densely or sparsely punctate; punctures on scutum uniformely distributed, seldom some interspaces amongpunctures larger than others [fig. 76]; T2 with fine, but deep and well distinct punctures, punctated area usually does not reachapical depression [fig. 82]; paraocular area above clypeus densely punctate, interspaces among large punctures small, filled bysmall punctures; gena narrow, but just behind eyes converging just slightly, in dorsal view seems to be angle between posterioreye margin and lateral margin of gena more than 45°; pygidial plate longer, in most specimens parallel-sided, apical margin ill-defined; glabrous base of T6 slightly wider than pygidium [fig. 86] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . nomioidis Pesenko

32 b. Frons in front of the oceli sparsely punctate, lateral parts of frons in some specimens densely punctate [fig. 75]; punctures onscutum uniformely distributed, except around paranotal ridges, where are distances often larger (like missing punctures); T2with well developed and deep punctures, punctated area usually reach apical depression in the middle of T2 and joiningpunctures on its margin [fig. 81]; paraocular area above clypeus more sparsely punctate, than in the rest of paraocular area, thisarea looks shiny almost like clypeus; gena narrow, just behind eyes strongly converging, in dorsal view seems to be anglebetween posterior eye margin and lateral margin of gena less than 45°; pygidial plate shorter, in most specimens narrowingapically, apical margin thick; glabrous base of T6 distinctly wider than pygidium [fig. 85] . . . . . . . . . . . . marginatus Hagens

Males ♂♂

1 a. Gonocoxite without impression; small to big species [fig. 139]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 b. Gonocoxite with impression; smaller species [fig. 159]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 222 a. Upper side of hind tibiae with red spines in pale pubescence [fig. 87]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 b. Upper side of hind tibiae only with pale pubescence, without any spine [fig. 88] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43 a. Occiput with transverse ridge [fig. cf. 2]; T4 and T5 finely and densely punctate; large species, gonostylus [figs. 197, 198], 9–

11 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . spinulosus Hagens3 b. Occiput without transverse ridge; T4 and T5 finely, very sparsely punctate with fine sculpture among punctures; medium size

species, gonostylus [figs. 163, 164], 6–8 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . majalis Pérez4 a. Felt-like pubescence covers larger part of flagellomere, flagellomeres only with small circular glabrous areas; flagellomere 2

short, maximally one and half longer than wide [fig. 90]; lower parts of mesopleurae with elongate ridges [fig. 89]; genitalusually pale brown with dark apex, gonostylus [figs. 191, 192], 6–8 mm . . . . . . . . . . . . . . . . . . . . . . . . . . rufiventris (Panzer)

4 b. Felt-like pubescence different; flagellomere 2 twice or more longer than wide; mesopleurae with grainy sculpture . . . . . . 105 a. Hind wing with 7–14 hamuli on the anterior part [fig. cf. 4]; upper and lower halves of flagellomeres conspicuously separated

by a ledge [fig. 95]; angle between cubital and anal veins on hind wing acute [fig. cf. 6] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65 b. Hind wing with 5–6 hamuli [fig. cf. 5] (only at some specimens of S. pellucidus, S. rubicundus and S. ruficrus up to 8 hamuli);

flagellomeres undivided [fig. 121]; angle between cubital and anal veins on hind wing acute to right. . . . . . . . . . . . . . . . . . 156 a. Sharp, crested ridge on gena [fig. cf. 8] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7



6 b. Without any ridge on gena . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97 a. Scutum densely and coarsely punctate with indistinct interspaces among punctures [fig. 93]; top of vertex with indistinct

longitudinal crista, gonostylus [figs. 193, 194], 9–12 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . scabricollis Wesmael7 b. Interspaces among punctures on scutum well developed; top of vertex with distinct longitudinal crista . . . . . . . . . . . . . . . . . . 88 a. Scutum sparsely, finely punctate with large interspaces [fig. 91]; T1 finely and sparsely punctate; felt-like pubescence on

flagellomeres 5–10 distinct, covers about 1/3 of the length of flagellomere [fig. 94]; sparse hair on head and thorax, gonostylus[fig. 149, 150], 6–8 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . cristatus Hagens

8 b. Scutum coarsely punctate [fig. 92]; T1 uniformely densely and coarsely punctate; felt-like pubescence on flagellomeres verynarrow, less than 1/4 of the length [fig. 95]; dense white hair on head and thorax, gonostylus [fig. 175, 176], 7–11 mm . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . olivieri Lepeletier

9 a. Scutum and tergites coarsely, densely punctate with minute interspaces among punctures [fig. 96]; the whole body looks dull,gonostylus [figs. 139, 140], 11–15 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . albilabris (Fabricius)

9 b. Scutum and tergites sparsely punctate with large interspaces among punctures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1010 a. Head about as long as wide [fig. 97]; apical parts of T1 to T3 very sparsely punctate [fig. 99]; gonostylus short and without

long hair [figs. 169, 170], 7–10 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . monilicornis (Kirby)10 b. Head wider than long [fig. 98]; apical parts of T1 and T2 more densely punctate; gonostylus longer and hairy . . . . . . . . . . . 1111 a. Vertex gibbous, behind ocelli strongly elevated, coarsely punctate with shiny interspaces [fig. 100]; apical depression of T2 to

T4 well defined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1211 b. Vertex behind ocelli slightly elevated, finely punctate and ridged [fig. 101]; apical depression of T2 to T4 ill defined . . . . . 1312 a. Flagellomeres twice longer than wide; felt-like pubescence on flagellomeres 5 to 10 overall base of flagellomeres [fig. 103];

apical depression of T4 smooth, unpunctate; gonostylus with elongated hairy projection [figs. 157, 158], 7–13 mm . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . gibbus (Linnaeus)

12 b. Flagellomeres less than twice as long as wide; felt-like pubescence on flagellomeres 5 to 10 short [fig. 102]; apical depressionof T4 with sparse coarse punctures; gonostylus projection not elongated [figs. 195, 196], 8–11 mm . . . . . . . schenckii Hagens

13 a. T4 finely punctate with microsculptured apical depression [fig. 104], gonostylus [fig. 185, 186], 7–10 mm . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . reticulatus Thomson

13 b. T4 coarsely punctate with smooth, shiny apical depression . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1414 a. Flagellomere 10 (forelast flagellomere) a little longer than wide [fig. 107]; T4 with well visible interspaces among punctures

[fig. 105], gonostylus [figs. 141, 142], 8–11 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . alternatus Smith14 b. Flagellomere 10 about 1.5x longer than wide [fig. 108]; T4 riddle-like punctate with indistinct interspaces [fig. 106],

gonostylus [figs. 143, 144], 8–11 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . crassanus Warncke15 a. The whole body (including legs and abdomen) black; hypoepimeral area smooth and shiny [fig. cf. 35]; head behind eyes

nearly as wide as compound eye, gonostylus [figs. 203, 204], 4.5–5.5 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . niger Hagens 15 b. The body always black and red, if only black, then tarsi reddish/yellow and hypoepimeral area punctate or sculptured . . . . 1616 a. T2 in basal half densely and coarsely punctate; apical part of T1 coarsely punctate [fig. 109]; gonostylus narrow, loaflike [figs.

187, 188] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1716 b. T2 sparsely punctate; apical part of T1 shiny with only few punctures [fig. 110]; frons with distinct interspaces among

punctures; gonostylus of another shape . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1817 a. Right angle between cubital and anal veins on hindwing [fig. cf. 44]; T4 dark in the middle, T5 completely dark, gonostylus

[figs. 187, 188], 8–12 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . rubicundus Hagens17 b. Acute angle between the cubital and anal veins on hindwing; T4 all, T5 reddish on sides; T1 often with basal dark spot [fig.

111], gonostylus [figs. 189, 190], 6–9 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ruficrus Erichson18 a. Felt-like pubescence on flagellomeres 8 to 10 short, up to 1/3 of the length of flagellomere [fig. 115]; T2 coarsely and densely

punctate [fig. 112]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1918 b. Felt-like pubescence on flagellomeres 8 to 10 long, at least 1/2 of the length of flagellomere [fig. 114]; T2 sparsely punctate

[fig. 113] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2019 a. Flagellomere 1 as wide as long [fig. 116]; gonostylus elongated and prominent [figs. 201, 202], 5–6 mm . ..dusmeti Blüthgen19 b. Flagellomere 1 distinctly wider than long [fig. 115]; gonostylus short [figs. 183, 184], 4.5–7 mm . . . . . puncticeps Thomson20 a. Head distinctly wider than thorax, with whitish long pubescence; metatarsus dark; felt-like pubescence on flagellomeres more

than ½ of the flagellomere [fig. 114]; wings hyaline, gonostylus [figs. 177, 178], 7–11 mm . . . . . . . . . . pellucidus Thomson20 b. Head not distincly wider than thorax and without long hair; metatarsus reddish; felt-like pubescence on flagellomeres about ½

of the flagellomere; wings slightly infumate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2121 a. Frons and vertex coarsely punctate, interspaces dull; scutellum densely punctate, interspaces smaller than punctures [fig. 117];

head distinctly wider than long, gonostylus [figs. 151, 152], 6–9 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . ephippius (Linnaeus)21 b. Frons and vertex sparsely and finely punctate, interspaces shiny; scutellum with few large punctures and large interspaces

among them [fig. 118]; head about as long as wide, gonostylus [figs. 171, 172], 3.5–5 mm. . . . . . . . . . . . . . longulus Hagens 22 a. Pronotal projections round [fig. cf. 48]; abdomen as wide as thorax or wider; face finely rugose, dull. . . . . . . . . . . . . . . . . . . 2322 b. Pronotal projections sharp [fig. cf. 49] (less distinct in some geoffrellus); abdomen narrower than thorax; face punctate, shiny

or dull . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2423 a. Tibiae and tarsi reddish; felt-like pubescence about or more than half of the length of the flagellomere [fig. 119]; end of

gonostylus curved [figs. 159, 160] , 5–7 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . hyalinatus Hagens 23 b. Tibiae and tarsi dark or black; felt-like pubescence as ¼ of the maximal length of flagellomere [fig. 120]; end of gonostylus

straight, triangular [figs. 153, 154], 6–9 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ferruginatus Hagens 24 a. Felt-like pubescence of middle flagellomeres not longer than half of flagellomere [fig. 121]. . . . . . . . . . . . . . . . . . . . . . . . . 25



24 b. Felt-like pubescence of middle flagellomeres distinctly longer than half of flagellomere, in some species cover all length offlagellomeres 4 to 11[figs. 122, 123] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27

25 a. Head a little longer than wide [fig. 124]; scutum with coarse but shallow punctures [fig. 129]; T1 and T2 with very fine, butnumerous punctures, gonostylus [figs. 181, 182], 5–8 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pseudofasciatus Blüthgen

25 b. Head round or wider than long [fig. 125]; T1 and T2 with fine and strong punctures mixed . . . . . . . . . . . . . . . . . . . . . . . . . . 2626 a. T1 with a few variable punctures [fig. 127]; gonostylus with large membranous part; apex of gonobase on ventral side with

patch of long hair [figs. 145, 146], 5–7 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . crassus Thomson26 b. T1 with numerous well-developed punctures [fig. 128]; gonostylus with small triangular membranous part; apex of gonobase

on ventral side with indistinct short hair [figs. 147, 148], 6–8 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . croaticus Meyer27 a. Frons with apressed white pubescence bellow as well as above antennal sockets; flagellomeres 2 to 11 long, more than 1.5x

longer than wide, covered by felt-like pubescence to the end of each segment [fig. 132] . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2827 b. Frons with apressed white pubescence bellow antennal sockets only; at least flagellomere 2 different . . . . . . . . . . . . . . . . . . 2928 a. Scutum densely punctate [fig. 131]; first abdominal tergites in most cases red, gonostylus [figs. 161, 162], 5–8 mm. . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . intermedius Blüthgen28 b. Scutum sparsely punctate [fig. 130]; abdomen usually dark to black, gonostylus [figs. 179, 180], 5–7 mm . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pinguiculus Pérez 29 a. T1 dorsally and T2 and T3 basally finely and very sparsely punctate, punctures nearly indistinct [fig. 133]; gonostylus short

[figs. 155, 156], 4–6 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . geoffrellus (Kirby)29 b. T2 and T3 basally densely punctate, punctures well-defined [fig. 134]; gonostylus short or long . . . . . . . . . . . . . . . . . . . . . 30 30 a. Interspaces among punctures on area between eye and lateral ocellus, as well as in front of the middle ocellus densely

microsculptured, dull [fig. 135]; T1 densely punctate [fig. 136]; scutum densely punctate, punctures deep and coarse [fig. 138].. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31

30 b. Interspaces among punctures on area between eye and lateral ocellus as well as in front of the middle ocellus shiny [fig. 137];T1 sparsely punctate; scutum coarsely, but sparsely punctate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32

31 a. Lower gena densely hairy, hair uniform in length, surface distinctly ridged and microsculptured, dull; vertex wide and long,upper gena behind eyes narrowing slowly, distance between hind ocellus and posterior margin of vertex almost as long asdistance between hind ocelli, vertex behind oceli flat; front ocellus always larger than hind ocelli; area of felt like pubescenceon flagellomeres strongly impressed [fig. 135], gonostylus [figs. 199, 200], 5–7 mm . . . . . . . . . . . . . . . zangherii Noskiewicz

31 b. Lower gena sparsely hairy, with intermixed long and short hair, surface distinctly microsculptured, but shiny; vertex narowerand shorter, upper gena behind eyes strongly narrowing, distance between hind ocellus and posterior margin of vertexdistinctly shorter than distance between hind ocelli, vertex behind ocelli decreasing; felt like pubescence on flagelomers veryvariable, gonostylus [figs. 147, 148], 6–8 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . croaticus Meyer

32 a. Membranous part of gonostylus enormous, trapezoidal; apex of gonobase on ventral side angulated, with patch of long hair[figs. 167, 168]; T1 very finely and sparsely punctate [fig. 134]; frons densely punctate, interspaces among punctures verysmall [fig. 126], 4–6 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . miniatus Hagens

32 b. Membranous part of gonostylus smaller; apex of gonobase on dorsal side with much shorter hair; T1 distinctly and denselypunctate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33

33 a. Membranous part of gonostylus large, slightly, but distinctly emarginated on its inner margin; apex of gonobase on ventral sideangulated, with patch of short, but distinct erected hair [fig. 173, 174]; head wider than long (distinct only in large specimens);felt-like pubescence on flagellomeres allways very long, from flagellomere 4 to 11 covers all length of flagellomere [fig. 123],3–6 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . nomioidis Pesenko

33 b. Membranous part of gonostylus small, triangular, on its inner margin straight; apex of gonobase on ventral side straight,without distinct erected hair [figs. 165, 166]; head round (distinct only in large specimens); felt-like pubescence offlagellomeres 3 to 6 very variable, but usually does not reach end of flagellomere, separated by elevated curled edge on distalmargin in most central-European specimens [fig. 122], 3–5 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . marginatus Hagens

List of species

Sphecodes albilabris (Fabricius, 1793)

Described as: Nomada albilabris Fabricius, 1793: 349. Synonyms: Dichroa fuscipennis Germar, 1819: Tab. 18.

Sphecodes latreillii Wesmael, 1835: 285.Sphecodes rubripes Spinola, 1838: 512.Sphecodes africanus Lepeletier, 1841: 541. Sphecodes nigripes Lepeletier, 1841: 542. Sphecodes rugosus Smith, 1848: 2209–2210. Sphecodes nodicornis Gistel, 1857: 554. Sphecodes fuscipennis var. basalis Dalla Torre, 1877: 185.Sabulicola cirsii Verhoeff, 1890: 329–331.Sphecodes grandis Meyer, 1922: 173.



Sphecodes rufipennis Cockerell, 1931: 348.Sphecodes atrescens Cockerell, 1931: 350.

Distribution. South and central Europe, in the north reaching Denmark, Estonia, southern Finland and Sweden,absent from Norway and on the British Isles; the distribution area extends to Asia (Westrich 1989, Warncke 1992,Lönnell & Cederberg 2007).

Biology. Species found in sandy sites, sand dunes, river banks and semideserts. Usually, it occurs in warmerregions and is locally highly abundant. Colletes cunicularius (Linnaeus) is the main host of this specialized cuckoobee (e.g. Blüthgen 1934, Westrich 1989). This cuckoo bee has only one generation during the year. It is highlyprobable that females survive for long time and fly in early summer, when Colletes is still not available. In this sit-uation S. albilabris can accept another (secondary) host. Old worn females were observed entering nests of Melit-turga clavicornis (Latreille) and larvae of S. albilabris were excavated and described from its nest by Rozen(1965). Blüthgen (1934) also stated Halictus quadricinctus (Fabricius) as unconfirmed host. We observed femalesof S. albilabris invading nests of this species at the exactly same situation as was Rozen’s (1965) observation ofthis species parasitizing Melliturga. Our record was made at Stroupeč Natural Monument in the Czech Republic inJuly 2011. However, suggested parasitic activity of the second generation, e.g. in nests of Dasypoda hirtipes (Fabr-icius), seems to be highly unlikely (A. Přidal, pers. comm.). Adult females and males emerge in July, mate, femalesfind shelter and overwinter. There is no place for parasitic activity before overwintering.

Taxonomic note. Warncke (1992) regarded S. rubripes Spinola, 1838 (with synonyms S. africanus Lepeletier,1841, Sphecodes rufipennis Cockerell, 1931, Sphecodes atrescens Cockerell, 1931) as subspecies of S. albilabris.However, the difference among S. albilabris, S. rubripes and S. africanus is major (Blüthgen, 1924) and not only incoloration as Warncke (1992) suggested. They also differ in phenology: males of S. rubripes and S. africanus werecollected in spring (April) and males of S. albilabris in summer. These forms can be valid species, but it requiresfurther taxonomic study.

Sphecodes alternatus Smith, 1853

Descibed as: Sphecodes alternatus Smith, 1853: 36.Synonyms: Sphecodes gibbus var. similis subvar. scariosus Sichel, 1865: 444.

Sphecodes punctiventris Hagens, 1882: 219.Sphecodes gracilior Morawitz, 1894: 78. Sphecodes antigae Tournier, 1901: 258–260.Sphecodes reticulatus var. algeriensis Alfken, 1914: 195.Sphecodes alternatus lindbergi Pittioni, 1950: 61–62.

Subspecies: S. a. gracilior Morawitz, 1894: 78. S. a. algeriensis Alfken, 1914: 195.

Distribution. South of Europe, reaching central Europe to Switzerland, Slovakia, Hungary, and Ukraine, in theeast through Turkey to Turkestan, present also in North Africa (Warncke 1992, Amiet et al. 1999).

Biology. Species of warm sites, especially sand dunes; quite common in various warm biotopes in southernEurope. In central Europe it is rare. Abundant in extensive sandy sites such as military training areas or larger sandquarries. Hosts unconfirmed. It was observed in association with Halictus compressus (Walckener) in Hungary(own observations); H. langobardicus Blüthgen is another likely host. Blüthgen (1934) recorded H. patellatus F.Morawitz as the likely host.

Sphecodes crassanus Warncke, 1992

Described as: Sphecodes crassanus Warncke, 1992: 26–27.No synonyms.

Distribution. South Europe—Spain, France (Warncke 1992), Switzerland (Amiet et al. 1999), Algeria.Biology. Poorly known species of open warm habitats with unknown host. The biology is probably similar to

S. alternatus and hosts can be expected within the medium sized species of Halictus.



Sphecodes crassus Thomson, 1870

Described as: Sphecodes crassus Thomson, 1870: 100.Synonyms: Sphecodes variegatus Hagens, 1874: 40–41.

Sphecodes divisus Hagens, 1882 (nec Kirby, 1802): 223. Sphecodes valesianus Frey-Gessner, 1903: 100.

Distribution. Europe north to 64° N, Sweden, Finland and Norway, present on British Isles; Turkey, Iran, andNorth Africa (Warncke 1992).

Biology. Common species that usually occurs in semi-open biotopes with shrubs, forest-steppes, steppes, andforest margins. In central Europe widespread, although except for warm sites it is never numerous. This species is anest cleptoparasite of smaller species of Halictidae. Sick et al. (1994) experimentally proved Stoeckhert’s (1933)finding of Lasioglossum pauxillum (Schenck) as a host, Westrich (1989) and Vegter (1993) mentioned also L.punctatissimum (Schenck) as a confirmed host. Other similar species are likely to be hosts as well: L.quadrinotatulum (Schenck) recorded as likely host by Alfken (1912), L. nitidiusculum (Kirby) by Stoeckhert(1933) and L. prasinum (Smith) by Vegter (1993). The hosts can be different in parts of the distributional area, as L.prasinum is rare in central but quite common in north-western Europe.

Sphecodes cristatus Hagens, 1882

Described as: Sphecodes cristatus Hagens, 1882: 218. No synonyms.

Distribution. Most of Europe north to 54°N, present in Sweden, absent on British Isles and Balkan Peninsula.Known from Turkey, Tajikistan and Mongolia (Westrich 1989, Warncke 1992).

Biology. Very rare species of sand dunes and larger sandy sites, in most countries known only from fewrecords. Blüthgen (1934) mentioned Lasioglossum nigripes (Lepeletier) as the likely host, but this species differsslightly in ecology. Westrich (1989) mentioned Halictus confusus Smith and H. subauratus (Rossi) as likely hosts,without any evidence. In our observations, the most likely host is H. leucaheneus Ebmer and probably also H.seladonius (Fabricius). We collected few S. cristatus at places with common occurence of H. leucaheneus inSlovakia, Hungary and Mongolia and the association seems to be possible. However, the most likely host and thecleptoparasite are both rare in central Europe, and to prove the host association is difficult.

Sphecodes croaticus Meyer, 1922

Described as: Sphecodes croaticus Meyer, 1922: 171.No synonyms.

Distribution. South Europe, reaching central Europe, known also from Cyprus, Russia, Turkey and Morocco(Warncke 1992). Due the confusion with S. pseudofasciatus and S. zangherii, the distribution can be incorrect. Weexamined material from: Austria, Bulgaria, Croatia, Czech Republic, France, Germany, Greece, Hungary, Italy,Portugal, Slovakia, Spain and Turkey.

Biology. Rare species of warm open habitats, usually steppes and warm sites, either on loess walls, or sandydunes; very rare and only in the warmest parts of central Europe. Lasioglossum interruptum (Panzer) is the onlypublished host (Blüthgen 1934) and was also observed by us and M. Herrmann (pers. comm.).

Note. Burger & Reum (2004) and Burger et al. (2006) stated that specimens from central Europe are not S.croaticus but instead S. zangherii. Examination of specimens from various parts of Europe and also specimensidentified by F. Burger showed that these specimens are in fact S. croaticus. Thus, S. croaticus is thermophilous andwidespread in central Europe, contrary to the much rarer S. zangherii.



Sphecodes dusmeti Blüthgen, 1924

Described as: Sphecodes dusmeti Blüthgen, 1924: 470–472.No synonyms.

Distribution. Southern Europe (Spain, France, Greece, Switzerland), North Africa (Morocco, Algeria) and Turkey(Warncke 1992).

Biology. Unknown.

Sphecodes ephippius (Linné, 1767)

Described as: Sphex ephippia Linné, 1767: 944.Synonyms: Apis minimus Harris, 1776 (nec Poda, 1761): Taf. 39, Fig. 21.

Apis obscura Geoffroy, 1785 (nec Linné, 1774, nec Müller, 1776): 447–448. Apis rufescens Geoffroy, 1785 (nec Gmelin, 1790): 447. Apis rufescens Gmelin, 1790 (nec Geoffroy, 1785): 2790. Apis labiata Fabricius, 1793: 342. Melitta divisa Kirby, 1802: 49–50. Andrena minuta Fabricius, 1804: 327. Sphecodes similis Wesmael, 1835: 283.Sphecodes zablocki Blüthgen, 1923c: 188.

Distribution. Europe to 62°N, present in Britain and Scandinavia, North Africa (Morocco to Egypt), Turkey, thedistribution area continues through Asia to Japan (Warncke 1992).

Biology. Dominant species in central and common in south Europe, where it is not as common probably due tocompetition with other species of the genus (S. gibbus, S. nomioidis). No biotope specialization; occurs nearlyeverywhere. Generalist with 18 known hosts, Bogusch et al. (2006) recorded the following species as confirmedhosts with complete references: Halictus tumulorum (Linnaeus), Lasioglossum laticeps (Schenck), L. leucozonium(Schrank), L. malachurum (Kirby), L. pauxillum and L. quadrinotatulum. In addition Andrena barbilabris (Kirby),A. flavipes Panzer, A. chrysopyga Schenck, A. labialis (Kirby), A. minutula (Kirby), A. wilkella (Kirby), Halictusmaculatus Smith, H. rubicundus (Christ), Lasioglossum fratellum (Péréz), L. lativentre (Schenck), and L. prasinumwere published as likely, unconfirmed hosts. The same authors also proved the individual specialization of thefemales of this species. It could be one of the reasons why is this species so numerous and widespread. Addition-ally, Andrena argentata Smith was observed as a host of S. ephippius.

Sphecodes ferruginatus Hagens, 1882

Described as: Sphecodes ferruginatus Hagens, 1882: 221.Synonym: Sphecodes rufescens var. alpestris Frey-Gessner, 1903: 107.

Distribution. Europe to 66°N, present in Scandinavia and Britain, Turkey, absent from North Africa (Warncke1992).

Biology. Species of midlands, usually bound to grasslands or sunny slopes, but not in warm areas. In centralEurope widespread but rare and uncommon. Lasioglossum fulvicorne (Kirby) is the only known documented host(Stoeckhert, 1933). Similar species such as L. laticeps and L. pauxillum have been published as likely hosts of thisspecies (Stoeckhert 1933, Westrich 1989). Surveys on localities with these species present did not support this asthe females of S. ferruginatus were invading only nests of L. fulvicorne.

Sphecodes geoffrellus (Kirby, 1802)

Described as: Melitta geoffrella Kirby, 1802: 45–46.Synonyms: Sphecodes affinis Hagens, 1882: 224.



Sphecodes fasciatus Hagens, 1882: 224. Sphecodes rimalis Pérez, 1903: CCXXI.Sphecodes impunctatus Meyer, 1922: 171–172.

Subspecies: Sphecodes geoffrellus atlasa Warncke, 1992: 36.Sphecodes geoffrellus hakkariensis Warncke, 1992: 36.

Distribution. Most of Europe, present in Britain and Scandinavia, North Africa (Morocco, Tunisia), Turkey, NearEast (Warncke 1992). Very numerous in southern Europe.

Biology. Common species both in warmer and cooler regions, with no strict biotope preferences. In centralEurope it is widespread and numerous, although recently sparser, probably due to the spread of similar speciesS. miniatus and S. nomioidis. Generalist parasitizing several hosts: Westrich (1989) has reported the followinghosts as confirmed Lasioglossum leucopus (Kirby), L. morio (Fabricius) and L. nitidiusculum, Bogusch (2003) L.pauxillum. Likely hosts are the following: L. fratellum published by Field (1996), L. rufitarse (Zetterstedt) byNeumeyer & Obrist (2000), L. sexstrigatum (Schenck) by Vegter (1993) and L. marginellum (Schenck) by Westrich(2006). It is possible that this species invades nests of various smaller Lasioglossum species, and the females areindividually specialized, as Bogusch et al. (2006) have confirmed in S. ephippius and S. monilicornis.

Sphecodes gibbus (Linnaeus, 1758)

Described as: Sphex gibba Linnaeus, 1758: 571. Synonyms: Apis glabra Füessly, 1775: 51.

Andrena ferruginea Olivier, 1789, nomen novum for Nomada gibba Fabricius, 1775 (nec Linnaeus, 1758): 139. Apis gibbosa Christ, 1791, nomen novum for Nomada gibba Fabricius, 1775 (nec Linnaeus, 1758): 177.Melitta picea Kirby, 1802: 48–49.Melitta sphecoides Kirby, 1802: 46–47. Andrena austriaca Fabricius, 1804: 325. Dichroa analis Illiger, 1806, nomen novum for Nomada gibba Fabricius, 1775 (nec Linnaeus, 1758): 48. Sphecodes apicatus Smith, 1853: 36–37.Sphecodes nigripennis Morawitz, 1876: 257. Sphecodes sutor Nurse, 1903: 538. Sphecodes gibbus var. rufispinosus Meyer, 1920: 113. Sphecodes gibbus var. turcestanicus Meyer, 1920: 113.Sphecodes nippon Meyer, 1922: 171.Sphecodes castilianus Blüthgen, 1924: 473–475. Sphecodes lustrans Cockerell, 1931: 411.Sphecodes pergibbus Blüthgen, 1938: 50.

Distribution. Nearly all of Europe (north to 63° N), the distribution area goes far into Asian mainland; also presentin North Africa (Warncke 1992).

Biology. Species of warm open or bushy habitats, more common in southern Europe (one of the mostnumerous Sphecodes species). In central Europe it shows affinity to warm biotopes, and is one of the most commonspecies of Sphecodes. This species is very variable in appearance and also in host spectrum. Main hosts are largeror medium sized species of the genus Halictus. Westrich (1989) recorded H. quadricinctus, H. rubicundus, and H.sexcinctus (Fabricius), Bogusch (2003) confirmed H. simplex Blüthgen as a host and presented Lasioglossummalachurum as a likely host. Other records of likely hosts are difficult to accept: Andrena vaga Panzer publishedby Möschler (1938) and Colletes cunicularius published by Riemann (1987). We observed a nesting site of A. vagafor several years, but no attempt to enter the nest by this cuckoo bee was observed, even though the activity of S.gibbus was observed few times. Westrich (1989) also put Halictus maculatus as likely host, which can be accepted.The list of hosts is probably larger than recently known and individual specialization to various hosts across thedistribution area is likely.

Sphecodes hyalinatus Hagens, 1882

Described as: Sphecodes hyalinatus Hagens, 1882: 222. No synonyms.



Distribution. Most of the Europe to 68°N, rare in south Europe; absent from Turkey and North Africa (Westrich1989, Warncke 1992).

Biology. Species of sunny sites in cooler regions, usually in rocky steppes on non-basic substrate. In centralEurope local and usually rare. Two known hosts are Lasioglossum fratellum confirmed by Field (1996) and L. ful-vicorne first mentioned by Stoeckhert (1933) as the likely host and since confirmed by Westrich (1989). The biol-ogy of L. fratellum and S. hyalinatus was studied by Heide (1992) and Field (1996). The abundance of this speciesis usually bound to the abundance of hosts. We have observed association of S. hyalinatus with both of the men-tioned hosts.

Sphecodes intermedius Blüthgen, 1923

Described as: Sphecodes intermedius Blüthgen, 1923a: 500–502.Synonym: Sphecodes lactipennis Meyer, 1925: 7–8.

Distribution. Very rare species, known from the Czech Republic, Slovakia, Hungary, Spain, Russia, and Turkey,Caucasus, Kazakhstan, Turkmenistan, Tajikistan, and Algeria (Warncke 1992). In all countries only one to severalrecords are known, and most of them are very old. Recently, the species was recorded at Bratčice in south Moravia(Czech Republic) in 2011.

Biology. Biology is poorly known; probably a species of warm and dry biotopes. Hosts unknown, in our opin-ion Halictus kessleri Bramson can be the host of this species. This bee is present at sand walls in Bratčice, wherewe found two specimens in 2011.

Sphecodes longulus Hagens, 1882

Described as: Sphecodes longulus Hagens, 1882: 226.Synonyms: Sphecodes longulus var. epidus Hagens, 1882: 226.

Sphecodes nitidulus Hagens, 1882: 226.

Distribution. Europe, north to south Finland, Sweden and Denmark, south England, eastwards to Tajikistan(Westrich 1989, Warncke 1992, Sörensson et al. 2009, Madsen & Calabuig 2011).

Biology. Rare species of sandy sites, uncommon in the localities in most of European countries. This species isnest cleptoparasite of small species of Lasioglossum; L. minutissimum (Kirby) is a confirmed host (Alfken 1912).Vegter (1993) brought evidence of two additional hosts: L. leucopus and L. zonulum (Smith), which is possible.Other likely hosts are the following: L. lucidulum (Schenck) and L. sexstrigatum published by Vegter (1993) and L.morio mentioned by Westrich (1989). During our observations nests of L. punctatissimum and L. semilucens(Alfken) were also invaded by females of this species.

Sphecodes majalis Pérez, 1903

Described as: Sphecodes majalis Pérez, 1903: CCXIX.Synonyms: Sphecodes gracilior Pérez, 1903: CCXIX.

Sphecodes problematicus Schulz, 1906: 235.Sphecodes barbarus Blüthgen, 1923a: 497.

Distribution. Spain, France, Belgium, Germany, Switzerland, Austria, Italy, Slovenia, Czech Republic, Slovakia,Hungary, Russia, Algeria (Warncke 1992).

Biology. Very rare species of steppes and sunny sites, bound to its host Lasioglossum pallens (Brullé). Her-rmann et al. (2003) described the biology both of the host and the parasite. The cleptoparasite as its host occur inspring (both males and females), and are recordable only during few days or weeks from March to May. Thus itseems to be rarer than it is.



Sphecodes marginatus Hagens, 1882

Described as: Sphecodes marginatus Hagens, 1882: 223.Synonyms: Sphecodes atratus Hagens, 1882: 224.

Sphecodes nigritulus Hagens, 1882: 225. Sphecodes biskrensis Pérez, 1903: CCXXI.

Subspecies: S. m. larochei Warncke, 1992: 34.S. m. creticus Warncke, 1992: 35.

Distribution. Unclear because of past confusion with S. nomioidis and reported by Warncke (1992) as S. marginatusbiskrensis. We examined wide material of the S. marginatus group and this species seems to be atlanto-mediterranean(absent from eastern Europe, Scandinavia and British Isles). Occurrence of S. marginatus was confirmed in thefollowing countries: Belgium, France, Germany (up to north), Italy (Sicilia), Morocco, Portugal, Spain, Tunisia andas subspecies in Canary Islands and Crete, also published for Denmark (Madsen & Calabuig 2011).

Biology. Species of warm open biotopes, usually sandy sites. In central Europe bound to heath and sandpitsand other open sandy biotopes. Hosts unknown, Vegter (1993) mentioned Dufourea minuta Lepeletier as likelyhost. However, the biology of both species is quite different, so we cannot accept it. In our opinion, small species ofLasioglossum occurring in sandy localities will be the hosts, eg. L. lucidulum, L. punctatissimum, L. semilucens, L.sexstrigatum or L. sabulosum, but this needs more study.

Sphecodes miniatus Hagens, 1882

Described as: Sphecodes miniatus Hagens, 1882: 223. Synonyms: Sphecodes dimidiatus Hagens, 1882: 224.

Sphecodes murithianus Frey-Gessner, 1903: 100.Sphecodes pilicornis Meyer, 1922: 170.

Distribution. Most of Europe (north to south Sweden), present in southern England; rare in southern Europe(confirmed only in Pelopponnese), absent from North Africa (Warncke 1992).

Biology. Common species especially of warmer and sandy sites. Westrich (1989) mentioned Lasioglossumnitidiusculum as confirmed host and Lasioglossum morio as likely host, Vegter (1993) L. sexstrigatum andBogusch (2003) L. politum Schenck. We also observed females of this species in association of L. pauxillum and L.punctatissimum. However, females of S. miniatus are similar to S. marginatus, and other small Sphecodes, so studyof the specialization in the field is difficult.

Sphecodes monilicornis (Kirby, 1802)

Described as: Melitta monilicornis Kirby, 1802: 47–48.Synonyms: Sphecodes maculatus Lepeletier, 1841: 545.

Sphecodes subquadratus Smith, 1845: 1014.Sphecodes gibbus var. subquadratus subvar. incertus Sichel, 1865: 420. Sphecodes gibbus var. ephippium subvar. nigrescens Sichel, 1865: 427.Sphecodes gibbus var. ephippium subvar. testaceipes Sichel, 1865: 428.Sphecodes gibbus var. ephippium subvar. rufipes Sichel, 1865: 428.Sphecodes gibbus var. subquadratus subvar. dubius Sichel, 1865: 419.Sphecodes ruficrus Dalla Torre, 1896 (nec Erichson, 1835), nomen novum for S. rufipes Sichel, 1865: 9. Sphecodes hanuman Nurse, 1903: 538–539. Sphecodes smyrnaensis Meyer, 1920: 116.Sphecodes caucasicus Meyer, 1920: 124.Sphecodes quadratus Meyer, 1920: 129. Sphecodes cephalotes Meyer, 1920: 129. Sphecodes monilicornis var. nigerrima Blüthgen, 1927: 41.Sphecodes quadratus cephalotiformes Pittioni, 1950: 62.

Subspecies: Sphecodes monilicornis quadratus Meyer, 1920: 129.



Sphecodes monilicornis cephalotes Meyer, 1920: 129.Sphecodes monilicornis berberus Warncke, 1992: 22.

Distribution. Most of Europe, north to 64°N, in the east through Asia to Japan, present in North Africa (Westrich1989, Warncke 1992).

Biology. Common species, especially in warm habitats. Occurs mainly on steppes and sandy sites. It seems tobe generalist with many hosts. Bogusch et al. (2006) summarized all known hosts of this species. Confirmed hostsare the following: Halictus rubicundus, Lasioglossum albipes (Fabricius), L. calceatum (Scopoli), L. leucozonium,L. quadrinotatulum and L. zonulum. L. malachurum is another confirmed host by our research. Vegter (1993)brought data of possible parasitation in nests of L. prasinum, Bogusch (2003) of Andrena flavipes, Halictusmaculatus, H. tumulorum, Lasioglossum laticeps, L. pauxillum and L. villosulum (Kirby). Thus the number of hostsof this species is high and the females are individually specialized. Most of the hosts are primitively eusocial andfemales of S. monilicornis lay eggs either into their nests within the solitary phase in spring or summer workerphase. Females of S. monilicornis can kill the workers of a nest and than lay eggs in all cells including those sealedand those not fully supplied (A. Přidal, pers. comm.).

Sphecodes niger Hagens, 1874

Described as: Sphecodes niger Hagens, 1874: 43. Synonym: Sphecodes niger var. holomelaena Blüthgen, 1949: 79.

Distribution. Europe from north-east Spain to Ukraine, Turkey, absent from Scandinavia and North Africa(Westrich 1989, Warncke 1992).

Biology. Species of warm biotopes, where can be common. Usually, it is not very abundant but widespread.The only confirmed host is Lasioglossum morio; Alfken (1912) also thought L. lucidulum to be a host. However,this species occurs mainly in sandy sites, where S. niger is uncommon.

Sphecodes nomioidis Pesenko, 1979

Described as: Sphecodes nomioidis Pesenko, 1979: 136.No synonyms.

Distribution. Unclear due to confusion with S. marginatus. We examined wide material of S. marginatus groupand S. nomioidis seems to be ponto-mediterranean (absent from west Europe and North Africa) species. The occur-rence of S. nomioidis was confirmed in the following countries: Austria, Bulgaria, Czech Republic, Greece, Hun-gary, Jordan, Romania, Slovakia and Turkey.

Biology. Poorly known, because of confusion with S. marginatus. It occurs on warm sandy and loess biotopesin central Europe. The species is expanding northwards recently and getting more common in central Europe.

Taxonomic note. Sphecodes marginatus biskrensis Pérez, 1903 sensu Warncke (1992) was characterized bystrongly elongated felt-like pubescence on antennomeres. This character seems to be very variable and presentcommonly in S. marginatus as well as in S. nomioidis. The type of S. biskrensis comes from North Africa, where S.nomioidis does not occur. For this reason, S. nomioidis remains the only available name for species from easternEurope. The application of the name needs to be confirmed by study of the type material as well as examination ofa wider material from the Ukraine and other eastern European localities.

Sphecodes olivieri Lepeletier, 1825

Described as: S. olivieri Lepeletier, 1825: 448.Synonyms: Sphecodes collaris Spinola, 1843: 137–139.

Sphecodes hispanicus var. abyssinicus Sichel, 1865: 447.Sphecodes ruficornis Sichel, 1865: 440.



Sphecodes punctulatus Sichel, 1865: 443–444.Sphecodes subpunctulatus Sichel, 1865: 445–446.Sphecodes rufithorax Morawitz, 1876: 255–256.Sphecodes verticalis Hagens, 1882: 219.Sphecodes desertus Nurse, 1903: 540–541.Sphecodes chionospilus Cockerell, 1911: 217–218. Sphecodes chionospilus var. sanguinatus Cockerell, 1911: 217.Sphecodes tenuis Meyer, 1920: 121–122. Sphecodes olivieri var. niveatus Meyer, 1925: 4.

Distribution: Southern Europe from Spain to Turkey, North Africa (Morocco to Egypt), Israel, Caucasus(Warncke 1992) and Iran.

Biology. Southern species occurring in warm semidesert habitats. The hosts of this rare species are not wellknown. Blüthgen (1934) mentioned Lasioglossum aegyptiellum (Strand) and L. vagans (Smith) as likely hosts.

Sphecodes pellucidus Smith, 1845

Described as: Sphecodes pellucidus Smith, 1845: 1014. Synonyms: Sphecodes pilifrons Thomson, 1870: 99.

Sphecodes brevicornis Hagens, 1874: 39–40. Sphecodes volatilis Smith, 1879: 26. Sphecodes pellucidus var. algirus Alfken, 1914: 195.Sphecodes pellucidus var. hypridus Blüthgen, 1925: 516. Sphecodes pellucidus var. niveipennis Meyer, 1925: 7.

Distribution. Europe north to 66°N, present in southern Sweden and Finland, east parts of Asia, North Africa(Westrich 1989, Warncke 1992).

Biology. Species of sandy sites, sand dunes and semidesert biotopes. Locally common. Alfken (1913) con-firmed host Andrena barbilabris. Witt (1992) described the biology of this host and the cleptoparasite. Schönitzer& Klinksik (1990) confirmed also A. nycthemera Imhoff; Sick et al. (1994) Lasioglossum leucozonium as hosts ofS. pellucidus. Other bee species of sandy biotopes were usually published as likely hosts: Andrena argentata Smith,A. bicolor Fabricius, A. humilis Imhoff, A. ventralis Imhoff and A. wilkella; Vegter (1993) mentioned also Lasio-glossum prasinum as likely host. This species was probably common in the localities investigated by this author,because it was considered a likely host of many Sphecodes species. In our survey, S. pellucidus was usuallyobserved in association with A. barbilabris, and S. ephippius from the same localities invaded nests of A. argen-tata. In our opinion, S. pellucidus is specialized and other hosts are unlikely. To the contrary, we could not rejectsimilar behaviour in S. albilabris, where females try to attack nests of other species.

Sphecodes pinguiculus Pérez, 1903

Described as: Sphecodes pinguiculus Pérez, 1903: CCXX. Synonyms: Sphecodes sareptensis Meyer, 1922: 170–171.

Sphecodes excellens Meyer, 1922: 170.Sphecodes punctatissimus Meyer, 1922: 172.Sphecodes hungaricus Blüthgen, 1923a: 498–499. Sphecodes coelebs Blüthgen, 1923a: 505–506.Sphecodes consobrinus Blüthgen, 1923a: 507–508.Sphecodes persicus Blüthgen, 1925: 509–511.Sphecodes capverdensis Pesenko & La Roche, 2002: 72, 203 in Pauly et al. (2002), syn. nov.

Taxonomy. We studied material of this species from Mongolia, Hungary, the Mediterranean region and Cape Verdeislands. Variablity of this species is not large and for this reason we propose synonymization of S. capverdensiswith S. pinguiculus.



Distribution. Spain, Hungary, Slovakia, Italy (Sicilia), Russia, North Africa, Turkey, Mongolia (Warncke1992), Cape Verde.

Biology. Species of warm, usually sandy habitats. This species is widely distributed in desert and semidesertregions, where it can be quite abundant. This species is almost surely a parasite of Halictus lucidipennis Smith ofCape Verde Islands. It is the only available common bee at the sites with occurence of S. pinguiculus and of course,close association of both species has been observed in Cape Verde. Possibly, H. lucidipennis is main host in mostareas of its occurrence, because this Halictus is common species in desert and semidesert areas from Mongolia,through Mediterranean region to Cape Verde. We suggest H. smaragdulus Vachal to be possible host for centralEuropean region, where similar H. lucidipennis does not occur. Nobile & Turrisi (2004) described seven new spe-cies related to S. pinguiculus. However, the characters are poor and probably relate to the way of preservation orcollecting rather than true structural differences. Recently, the type material was studied (Schwarz & Gusenleitner2012) and the described species are not related to S. pinguiculus but S. miniatus group.

Sphecodes pseudofasciatus Blüthgen, 1925

Described as: Sphecodes pseudofasciatus Blüthgen, 1925: 473. No synonyms.

Distribution. Distribution is poorly known, because of former incorrect synonymization of this species under S.croaticus by Warncke (1992). We examined material from: Austria, Czech Republic, France, Hungary, Italy, Portu-gal, Romania, Russia, Slovakia, Spain, Switzerland, Morocco, Turkey and Ukraine.

Biology. This species is collected in low numbers and only few records are known from each country. Host isunknown.

Sphecodes puncticeps Thomson, 1870

Described as: Sphecodes puncticeps Thomson, 1870: 99–100. Synonyms: Sphecodes bituberculatus Pérez, 1903: CCXX–CCXXI.

Sphecodes opacifrons Pérez, 1903: CCXIX–CCXX.Sphecodes puncticeps var. cretanus Strand, 1921: 305.

Distribution. Europe north to south Finland and Sweden, Asia, North Africa to Egypt, Israel (Westrich 1989,Warncke 1992), Iran.

Biology. Species of warm sandy biotopes, in south Europe quite common, in central Europe local but in sandylocalities usually abundant. The only confirmed host is Lasioglossum villosulum, published by Alfken (1913). Bis-choff (1927) put L. brevicorne (Schenck) as likely host. We confirm both hosts and have observed S. puncticepsalso in an association with L. politum and L. sabulosum (Warncke), but with no evidence.

Sphecodes reticulatus Thomson, 1870

Described as: Sphecodes reticulatus Thomson, 1870: 98. Synonym: Sphecodes distinguendus Hagens, 1874: 38–39.

Distribution. Europe north to 62°N, present in south Sweden and Finland, British Isles, east to Turkestan. In theMediterranean very rare, known from Bulgaria, Greece and Turkey (Westrich 1989, Warncke 1992).

Biology. Species of various biotopes, usually in sandy sites, heathlands and sandpits. Present also in other openor semi-open habitats, as steppes, shrubby sites, etc. Rare in the whole distribution area. Andrena barbilabris wasmentioned as confirmed host by Stoeckhert (1933) and Blüthgen (1934); A. argentata, A. wilkella, Lasioglossumprasinum and L. leucozonium cedri Ebmer as likely hosts (Blüthgen 1934, Vegter 1993, M. Schwarz, pers. comm.).



Sphecodes rubicundus Hagens, 1875

Described as: Sphecodes rubicundus Hagens, 1875: 318. Synonym: Sphecodes rubicundus altisilesiacus Torka, 1926: 126–127.

Distribution. Europe north to 56° N, England, eastern to Ural and Armenia. Present in Turkey (Westrich 1989,Warncke 1992).

Biology. Very rare species. Older findings are from open habitats: usually sandy sites, sanddunes and steppes.Sowa & Mostowska (1978) described the biology of Andrena labialis (Kirby) and confirmed it as a host of this spe-cies. Blüthgen (1934) also put A. labialis as confirmed and A. nigroaenea (Kirby) as likely host of S. rubicundus.Torka (1925) brought data on possible parasitization in nests of A. agilissima (Scopoli). Currently, the mentionedhosts occur mostly in different biotopes from the cleptoparasite.

Sphecodes ruficrus (Erichson, 1835)

Described as: Dichroa ruficrus Erichson, 1835: 101–102.Synonyms: Sphecodes hispanicus Wesmael, 1835: 285–286.

Sphecodes rufipes Smith, 1853: 37.Sphecodes gibbus var. tunetanus Gribodo, 1894: 293–294. Sphecodes atrohirtus Pérez, 1903 (nomen novum for S. hispanicus Wesmael, 1836 sensu Hagens, 1882): CCXIX.

Distribution. South and western Europe: Spain, France, Switzerland, Italy, Germany, Hungary, Russia. This spe-cies extends its distribution area to north-east (Westrich 1989, Warncke 1992).

Biology. Species of sandy sites, in southern Europe common in such localities. Herrmann (2006) described thebiology and hosts. The main host is Andrena humilis, also related species A. livens Pérez and A. taraxaci Giraud arepossible hosts of this species. We have studied this species in Spain in association with A. livens. Westrich (1989)reported Andrena decipiens Schenck as a host, but with no further details.

Sphecodes rufiventris (Panzer, 1798)

Described as: Tiphia rufiventris Panzer, 1798: 4. Synonyms: Sphecodes subovalis Schenck, 1853: 223–224.

Sphecodes brevis Hagens, 1875: 317–318. Sphecodes singularis Meyer, 1920: 130. Sphecodes combinatus Blüthgen, 1927: 37–39.Sphecodes tadschicus Blüthgen, 1935: 366.Sphecodes subovalis austrinus Erlandsson, 1979: 123. Sphecodes subovalis austrinus var. balcanicus Erlandsson, 1979: 123.Sphecodes rufiventris hethiticus Warncke, 1992: 28.

Distribution. Europe up to 57°N, absent from Scandinavia and Britain, in the east the distribution area goes to UralRiver, present in North Africa: Morocco, Algeria (Warncke 1992).

Biology. Species of warmer open or shrubby habitats (forest steppes), common on loess walls. In centralEurope quite rare, more abundant only at xerothermes. The only known host is Halictus maculatus, recorded byStoeckhert (1933) and Blüthgen (1934). We have only observed S. rufiventris in association with this species.

Sphecodes scabricollis Wesmael, 1835

Described as: Sphecodes scabricollis Wesmael, 1835: 287.Synonym: Sphecodes perversus Ritsema, 1879: 56–57.



Distribution. Europe north to southern Finland, present in England, southward absent from the Balkan Peninsulaand Turkey (Westrich 1989, Warncke 1992).

Biology. Very rare species of sandy wetlands, in most countries endangered or extinct. The distribution is muchweaker than in the past. The host is Lasioglossum zonulum, recorded in the literature (Blüthgen 1934, Yates 2002),the first author also recorded Halictus compressus, H. quadricinctus and Lasioglossum prasinum as likely hosts.More recently, parasitation only in nests of L. zonulum was observed (M. Herrmann, pers. comm.).

Sphecodes schenckii Hagens, 1882

Described as: Sphecodes schenckii Hagens, 1882: 217–218. Synonyms: Sphecodes sulcicollis Pérez, 1903: CCVIII.Subspecies: Sphecodes schenckii caspicus Meyer, 1920: 113–114.

Distribution. North-east Spain, France, Switzerland, Italy, Hungary, Austria, Slovakia, Germany, Russia, Turkeyand Iran (Warncke 1992).

Biology. Rare species of warm localities, in most of the countries only few records. Lasioglossum discum(Smith) is supposed to be a host due to the similar distribution area, size and appearance (Blüthgen 1934). Groz-danić (1971) confirmed this species as host of S. schenckii. In northern Switzerland and in Germany L. discum ismissing, so there has to be one or more other hosts, probably Halictus simplex (M. Herrmann, pers. comm. andpers. obs.).

Sphecodes spinulosus Hagens, 1875

Described as: Sphecodes spinulosus Hagens, 1875: 317. No synonyms.

Distribution. Europe north to 56°N, present in England, Turkey, south-east Russia, and the distribution areaextends probably east to Asia. Present in North Africa (Warncke 1992, Edwards & Telfer 2001).

Biology. Rare species of warm biotopes, usually steppes. In central Europe only few records in recent years.The only known host is Lasioglossum xanthopus (Kirby) recorded by Stoeckhert (1933) and Blüthgen (1934).

Sphecodes zangherii Noskiewicz, 1931

Described as: Sphecodes zangherii Noskiewicz, 1931: 139.No synonyms.

Distribution. The distribution is poorly known due the taxonomical problems with this species in the past.Warncke (1992) suggested wide distribution in south and central Europe, from France to Turkey and north to Ger-many and Czech Republic. However, we examined large part of Warncke’s material and found only few specimensfrom the Alps in France and Switzerland, north and central Italy and mountains in Greece and Turkey. All otherspecimens of S. zangherii determined by Warncke were missidentified S. croaticus.

Biology. Rare species of warmer localities in mountains, but sometimes also in low altitudes. Details of itsbiology are unknown.



TABLE 1. List of hosts of Sphecodes Latreille species of central Europe. Likely hosts are marked by asterisk.

Species Hosts

Sphecodes albilabris (Fabricius, 1793) Colletes cunicularius (Linnaeus)

Halictus quadricinctus (Fabricius)

Melitturga clavicornis (Latreille)

*Dasypoda hirtipes (Fabricius)

Sphecodes alternatus Smith, 1853 *Halictus compressus (Walckener)

*Halictus langobardicus Blüthgen

*Halictus patellatus F. Morawitz

Sphecodes crassanus Warncke, 1992 hosts unknown

Sphecodes crassus Thomson, 1870 Lasioglossum pauxillum (Schenck)

Lasioglossum punctatissimum (Schenck)

*Lasioglossum quadrinotatulum (Schenck)

*Lasioglossum nitidiusculum (Kirby)

*Lasioglossum prasinum (Smith)

Sphecodes cristatus Hagens, 1882 *Halictus confusus Smith

*Halictus leucaheneus Ebmer

*Halictus seladonius (Fabricius)

*Halictus subauratus (Rossi)

*Lasioglossum nigripes (Lepeletier)

Sphecodes croaticus Meyer, 1922 *Lasioglossum interruptum (Panzer)

Sphecodes dusmeti Blüthgen, 1924 hosts unknown

Sphecodes ephippius (Linné, 1767) Andrena argentata Smith

Halictus tumulorum (Linnaeus)

Lasioglossum laticeps (Schenck)

Lasioglossum leucozonium (Schrank)

Lasioglossum malachurum (Kirby)

Lasioglossum pauxillum Schenck

Lasioglossum quadrinotatulum (Schenck)

*Andrena barbilabris (Kirby)

*Andrena flavipes Panzer

*Andrena chrysopyga Schenck

*Andrena labialis (Kirby)

*Andrena minutula (Kirby)

*Andrena wilkella (Kirby)

*Halictus maculatus Smith

*Halictus rubicundus (Christ)

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TABLE 1. (continued)

Species Hosts

Sphecodes ephippius (Linné, 1767) *Lasioglossum fratellum (Péréz)

*Lasioglossum lativentre (Schenck)


Sphecodes ferruginatus Hagens, 1882 Lasioglossum fulvicorne (Kirby)

*Lasioglossum laticeps (Schenck)

*Lasioglossum pauxillum Schenck

Sphecodes geoffrellus (Kirby, 1802) Lasioglossum leucopus (Kirby)

Lasioglossum morio (Fabricius)

Lasioglossum nitidiusculum (Kirby)

Lasioglossum pauxillum Schenck

*Lasioglossum fratellum (Péréz)

*Lasioglossum marginellum (Schenck)

*Lasioglossum rufitarse (Zetterstedt)

*Lasioglossum sexstrigatum (Schenck)

Sphecodes gibbus (Linnaeus, 1758) Halictus quadricinctus (Fabricius)

Halictus rubicundus (Christ)

Halictus sexcinctus (Fabricius)

Halictus simplex Blüthgen

*Andrena vaga Panzer

*Colletes cunicularius (Linnaeus)


*Lasioglossum malachurum (Kirby)

Sphecodes hyalinatus Hagens, 1882 Lasioglossum fratellum (Péréz)

Lasioglossum fulvicorne (Kirby)

Sphecodes intermedius Blüthgen, 1923 *Halictus kessleri Bramson

Sphecodes longulus Hagens, 1882 Lasioglossum leucopus (Kirby)

Lasioglossum minutissimum (Kirby)

Lasioglossum punctatissimum (Schenck)

Lasioglossum semilucens (Alfken)

Lasioglossum zonulum (Smith)

*Lasioglossum lucidulum (Schenck)

*Lasioglossum sexstrigatum

Sphecodes majalis Pérez, 1903 Lasioglossum pallens (Brullé)

Sphecodes marginatus Hagens, 1882 *Dufourea minuta Lepeletier

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Species Hosts

Sphecodes marginatus Hagens, 1882 * Lasioglossum lucidulum (Schenck)

*Lasioglossum punctatissimum (Schenck)

*Lasioglossum sabulosum (Warncke)

*Lasioglossum semilucens (Alfken)


Sphecodes miniatus Hagens, 1882 Lasioglossum nitidiusculum (Kirby)

*Lasioglossum morio (Fabricius)


*Lasioglossum politum Schenck

*Lasioglossum punctatissimum (Schenck)


Sphecodes monilicornis (Kirby, 1802) Halictus rubicundus (Christ)

Lasioglossum albipes (Fabricius)

Lasioglossum calceatum (Scopoli)


Lasioglossum malachurum (Kirby)

Lasioglossum quadrinotatulum (Schenck)

Lasioglossum zonulum (Smith)

*Andrena flavipes Panzer


*Halictus tumulorum (Linnaeus)

*Lasioglossum laticeps (Schenck)



*Lasioglossum villosulum (Kirby)

Sphecodes niger Hagens, 1874 Lasioglossum morio (Fabricius)

*Lasioglossum lucidulum (Schenck)

Sphecodes nomioidis Pesenko, 1979 hosts unknown

Sphecodes olivieri Lepeletier, 1825 *Lasioglossum aegyptiellum (Strand)

*Lasioglossum vagans (Smith)

Sphecodes pellucidus Smith, 1845 Andrena barbilabris (Kirby)

Andrena nycthemera Imhoff


*Andrena argentata Smith

*Andrena bicolor Fabricius

*Andrena humilis Imhoff

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Species Hosts

Sphecodes pellucidus Smith, 1845 *Andrena ventralis Imhoff



Sphecodes pinguiculus Pérez, 1903 Halictus lucidipennis Smith

*Halictus smaragdulus Vachal

Sphecodes pseudofasciatus Blüthgen, 1925 hosts unknown

Sphecodes puncticeps Thomson, 1870 Lasioglossum villosulum (Kirby)

*Lasioglossum brevicorne (Schenck

*Lasioglossum politum Schenck

*Lasioglossum sabulosum (Warncke)

Sphecodes reticulatus Thomson, 1870 Andrena barbilabris (Kirby)

*Andrena argentata Smith


*Lasioglossum leucozonium cedri Ebmer


Sphecodes rubicundus Hagens, 1875 Andrena labialis (Kirby)

*Andrena agilissima (Scopoli)

*Andrena nigroaenea (Kirby)

Sphecodes ruficrus (Erichson, 1835) Andrena humilis Imhoff

Andrena livens Pérez

*Andrena decipiens Schenck

*Andrena taraxaci Giraud

Sphecodes rufiventris (Panzer, 1798) Halictus maculatus Smith

Sphecodes scabricollis Wesmael, 1835 Lasioglossum zonulum (Smith)

*Halictus compressus (Walckener)

*Halictus quadricinctus (Fabricius)


Sphecodes schenckii Hagens, 1882 Lasioglossum discum (Smith)

*Halictus simplex Blüthgen

Sphecodes spinulosus Hagens, 1875 Lasioglossum xanthopus (Kirby)

Sphecodes zangherii Noskiewicz, 1931 hosts unknown



FIGURES 1–9. Females. 1—Sphecodes majalis, head frontal view, clypeus; 2—S. spinulosus, vertex with ridge; 3—S. spinu-losus, antenna; 4—S. schenckii, eight wing hammuli; 5—S. puncticeps, five wing hammuli; 6—S. crassanus, hindwing vena-tion; 7—S. hyalinatus, hindwing venation; 8—S. scabricolis, occiput with ridge; 9—S. scabricolis, head and scutum, dorsalview.



FIGURES 10–17. Females. 10—Sphecodes olivieri, dorsal view; 11—S. cristatus, vertex with longitudinal ridge; 12—S.monilicornis, head, frontal view; 13—S. albilabris, scutum; 14—S. monilicornis, mesopleura; 15—S. rufiventris, mesopleura;16—S. reticulatus, head, lateral view; 17—S. gibbus, head, lateral view.



FIGURES 18–27. Females. 18—Sphecodes reticulatus, tergum 4; 19—S. gibbus, tergum 4; 20—S. alternatus, tergum 4; 21—S. crassanus, tergum 4; 22—S. alternatus, head, frontal view; 23—S. crassanus, head, frontal view; 24—S. gibbus, scutum;25—S. schenckii, scutum; 26—S. gibbus, scapus; 27—S. schenckii, scapus.



FIGURES 28–38. Females. 28—Sphecodes pinguiculus, metasoma, sculpture of tergites 1–3; 29—S. pinguiculus, scutum;30—S. intermedius, scutum; 31—S. pinguiculus, head, frontal view; 32—S. puncticeps, head, frontal view; 33—S. crassus,head, frontal view; 34—S. longulus, head, frontal view; 35—S. niger, mesopleura; 36—S. niger, propodeum, dorsal view; 37—S. puncticeps, tergum 3; 38—S. longulus, tergum 3.



FIGURES 39–47. Females. 39—Sphecodes pellucidus, pygidium; 40—S. crassus, pygidium; 41—S. crassus, sculpture ofclypeus; 42—S. ephippius, sculpture of clypeus; 43—S. ruficrus, all body, dorsal view; 44—S. rubicundus, hind wing venation;45—S. rubicundus, apical margin of tergum 1; 46—S. ephippius, apical margin of tergum 1; 47—S. pellucidus, head, frontalview.



FIGURES 48–54. Females. 48—Sphecodes ferruginatus, pronotum, oblique view; 49—S. geoffrellus, pronotum, obliqueview; 50—S. hyalinatus, propodeum, dorsal view; 51—S. ferruginatus, thorax, ventral view; 52—S. hyalinatus, thorax, ventralview; 53—S. ferruginatus, head, frontal view; 54—S. hyalinatus, head, frontal view.



FIGURES 55–61. Females. 55—Sphecodes pseudofasciatus, head, frontal view; 56—S. pseudofasciatus, head, dorsal view;57—S. zangherii, head, frontal view; 58—S. zangherii, head, dorsal view; 59—S. croaticus, head, frontal view; 60—S. croati-cus, head, dorsal view; 61—S. croaticus, sculpture of tergite 2.



FIGURES 62–73. Females. 62—Sphecodes dusmeti, head, frontal view; 63—S. crassus, head, frontal view; 64—S. geoffrel-lus, head, frontal view; 65—S. crassus, dorsal part of mesopleura; 66—S. geoffrellus, dorsal part of mesopleura; 67—S. cras-sus, sculpture of tergum 2 and 3; 68—S. geoffrellus, sculpture of tergum 2 and 3; 69—S. crassus, antenna; 70—S. geoffrellus,antenna; 71—S. miniatus, antenna; 72—S. miniatus, head, lateral view; 73—S. nomioidis, head, lateral view.



FIGURES 74–86. Females. 74—Sphecodes miniatus, head, frontal view; 75—S. marginatus, head, frontal view; 76—S. nomi-oidis, head, frontal view; 77—S. miniatus, scutum; 78—S. marginatus, scutum; 79—S. nomioidis, scutum; 80—S. miniatus,sculpture of tergum 2; 81—S. marginatus, sculpture of tergum 2; 82—S. nomioidis, sculpture of tergum 2; 83, 84—S. miniatus,pygidium, oblique and dorsal view; 85—S. marginatus, pygidium, dorsal view; 86—S. nomioidis, pygidium, dorsal view.



FIGURES 87–96. Males. 87—Sphecodes spinulosus, hind tibia, lateral view; 88—S. monilicornis, hind tibia, lateral view;89—S. rufiventris, mesopleura; 90—S. rufiventris, antenna; 91—S. cristatus, scutum; 92—S. olivieri, scutum; 93—S. scabri-collis, head and scutum, dorsal view; 94—S. cristatus, antenna; 95—S. olivieri, antenna; 96—S. albilabris, scutum.



FIGURES 97–108. Males. 97—Sphecodes monilicornis, head, frontal view; 98—S. reticulatus, head, frontal view; 99—S.monilicornis, sculpture of tergites 1–3; 100—S. gibbus, head, sculpture of vertex; 101—S. reticulatus, head, sculpture of ver-tex; 102—S. schenckii, head, sculpture of vertex and antenna; 103—S. gibbus, antenna; 104—S. reticulatus, sculpture of tergite4; 105—S. alternatus, sculpture of tergite 4; 106—S. crassanus, sculpture of tergite 4; 107—S. alternatus, terminal flagellom-eres; 108—S. crassanus, terminal flagellomeres.



FIGURES 109–118. Males. 109—Sphecodes ruficrus, sculpture of tergites 1–2; 110—S. ephippius, sculpture of tergites 1–2;111—S. ruficrus, all body, dorsal view; 112—S. puncticeps, sculpture of tergites 1–2; 113—S. pellucidus, sculpture of tergites1–2; 114—S. pellucidus, antenna; 115—S. puncticeps, antenna; 116—S. dusmeti, antenna; 117—S. ephippius, sculpture ofscutellum; 118—S. longulus, sculpture of scutellum.



FIGURES 119–132. Males. 119—Sphecodes hyalinatus, antenna; 120—S. ferruginatus, antenna; 121—S. pseudofasciatus,antenna; 122—S. marginatus, antenna; 123—S. nomioidis, antenna; 124—S. pseudofasciatus, head, frontal view; 125—S. cras-sus, head, frontal view; 126—S. miniatus, head, frontal view; 127—S. crassus, sculpture of tergum 1; 128—S. croaticus, sculp-ture of tergum 1; 129—S. pseudofasciatus, scutum; 130—S. pinguiculus, scutum; 131—S. intermedius, scutum; 132—S.pinguiculus, head, frontal view.



FIGURES 133–138. Males. 133—Sphecodes geoffrellus, sculpture of tergites 1–3; 134—S. miniatus, sculpture of tergites 1–3;135—S. zangherii, head, dorsal oblique view; 136—S. zangherii, sculpture of tergum 1; 137—S. miniatus, head, dorsal obliqueview; 138—S. zangherii, scutum.



FIGURES 139–174. Male genitalia, dorsal and lateral views. 139, 140—Sphecodes albilabris; 141, 142—S. alternatus;143, 144—S. crassanus; 145, 146—S. crassus; 147, 148—S. croaticus; 149, 150—S. cristatus; 151, 152—S. ephippius; 153,154—S. ferruginatus; 155, 156—S. geoffrellus; 157, 158—S. gibbus; 159, 160—S. hyalinatus; 161, 162—S. intermedius; 163,164—S. majalis; 165, 166—S. marginatus; 167, 168—S. miniatus; 169, 170—S. monilicornis; 171, 172—S. longulus; 173,174—S. nomioidis.



FIGURES 175–204. Male genitalia, dorsal and lateral views. 175, 176—Sphecodes olivieri; 177, 178—S. pellucidus; 179,180—S. pinguiculus; 181, 182—S. pseudofasciatus; 183, 184—S. puncticeps; 185, 186—S. reticulatus; 187, 188—S. rubicun-dus; 189, 190—S. ruficrus; 191, 192—S. rufiventris; 193, 194—S. scabricollis; 195, 196—S. schenckii; 197, 198—S. spinulo-sus; 199, 200—S. zangherii; 201, 202—S. dusmeti; 203, 204—S. niger.

Acknowledgement

We would like to thank colleagues and collectors who helped us with the material. Special thanks belong to Maxi-milian Schwarz (Ansfelden, Austria) and Christian Schmid-Egger (Berlin, Germany), who helped us with materialfor this study and acccess to their collections. Additional thanks to Mike Herrmann (Konstanz, Germany) and one



anonymous referee for valuable comments to the article. Institutional support to JS was given by Ministry of Edu-cation of the Czech Republic, project no. MSM0021620828; grant support to PB and JS was given by Czech Sci-ence Foundation, project no. 206/09/0522.

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Review and identification of the cuckoo bees of central Europe (Hymenoptera: Halictidae: Sphecodes)

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